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vitellophags—traverse the yolk and assist in its rearrangement; he insists
on the importance both as regards quantity and quality of the yolk.
The eggs of some insects are fairly transparent, and the process of
development in them can, to a certain extent, be observed by simple
inspection with the microscope; a method that was used by Weismann in
his observations on the embryology of Chironomus. There is a moth
(Limacodes testudo), that has no objection to depositing its eggs on glass
microscope-slides. These eggs are about a millimetre long, somewhat
more than half that width, are very flat, and the egg-shell or chorion is
very thin and perfectly transparent. When first laid the contents of this egg
appear nearly homogeneous and evenly distributed, a finely granular
appearance being presented throughout; but in twenty-four hours a great
change is found to have taken place. The whole superficial contents of
the egg are at that time arranged in groups, having the appearance of
separate rounded or oval masses, pressed together so as to destroy
much of their globular symmetry. The egg contents are also divided into
very distinct forms, a granular matter, and a large number of transparent
globules, these latter being the fatty portion of the yolk; these are present
everywhere, though in the centre there is a space where they are very
scanty, and they also do not extend quite to the circumference. But the
most remarkable change that has taken place is the appearance in the
middle of the field of an area different from the rest in several particulars;
it occupies about one-third of the width and one-third of the length; it has
a whiter and more opaque appearance, and the fat globules in it are
fewer in number and more indistinct. This area is afterwards seen to be
occupied by the developing embryo, the outlines of which become
gradually more distinct. Fig. 83 gives an idea of the appearance of the
egg about the middle period of the development. In warm weather the
larva emerges from this egg ten or eleven days after it has been
deposited.
Fig. 83.—A, Egg of Limacodes testudo about the middle of the development
of the embryo; B, micropyles and surrounding sculpture of chorion.
Metamorphosis.
There are three great fields of inquiry in regard to metamorphosis, viz. (1)
the external form at the different stages; (2) the internal organs and their
changes; (3) the physiological processes. Of these only the first has yet
received any extensive attention, though it is the third that precedes or
underlies the other two, and is the most important. We will say a few
words about each of these departments of the inquiry. Taking first the
external form—the instar. But before turning to this we must point out that
in limiting the inquiry to the post-embryonic development, we are making
one of those limitations that give rise to much misconception, though they
are necessary for the acquisition of knowledge as to any complex set of
phenomena. If we assume five well-marked stages as constituting the life
of an Insect with extreme metamorphosis, viz. (1) the formation and
growth of the egg; (2) the changes in the egg culminating in its hatching
after fertilisation; (3) the period of growth; (4) the pupal changes; (5) the
life of the perfect Insect; and if we limit our inquiry about development to
the latter three, we are then shutting out of view a great preliminary
question, viz. whether some Insects leave the egg in a different stage of
development to others, and we are consequently exposing ourselves to
the risk of forgetting that some of the distinctions we observe in the
subsequent metamorphosis may be consequential on differences in the
embryonic development.
Instar and Stadium.
Every Insect after leaving the egg undergoes during the process of
growth castings of the skin, each of which is called a moult or ecdysis.
Taking for our present purpose five as the number of ecdyses undergone
by both the locust and butterfly, we may express the differences in the
successions of change we portray in Figs. 84 and 85 by saying that
previous to the first ecdysis the two Insects are moderately dissimilar, that
the locust undergoes a moderate change before reaching the fifth
ecdysis, and undergoes another moderate change at this moult, thus
reaching its perfect condition by a slight, rather gradual series of
alterations of form. On the other hand, the butterfly undergoes but little
modification, remaining much in the condition shown by A, Fig. 85, till the
fourth, or penultimate, ecdysis, but then suffers a complete change of
form and condition, which apparently is only inferior to another
astonishing change that takes place at the fifth or final moult. The chief,
though by no means the only, difference between the two series consists
in the fact that the butterfly has interposed between the penultimate and
the final ecdyses a completely quiescent helpless condition, in which it is
deprived of external organs of sense, locomotion, and nutrition; while in
the locust there is no loss of these organs, and such quiescent period as
exists is confined to a short period just at the fifth ecdysis. The changes
exhibited by the butterfly are called "complete metamorphosis," while this
phenomenon in the locust is said to be "incomplete." The Insect with
complete metamorphosis is in its early stage called a larva, and in the
quiescent state a pupa. The adult state in both butterfly and locust is
known as imago or perfect Insect.
The intervals between the ecdyses are called stadia, the first stadium
being the period between hatching and the first ecdysis. Unfortunately no
term is in general use to express the form of the Insect at the various
stadia; entomologists say, "the form assumed at the first moult," and so
on. To avoid this circumlocution it may be well to adopt a term suggested
by Fischer,[81] and call the Insect as it appears at hatching the first instar,
what it is as it emerges from the first ecdysis the second instar, and so on;
in that case the pupa of a Lepidopteron that assumed that condition at the
fifth ecdysis would be the sixth instar, and the butterfly itself would be the
seventh instar.
Various terms are used to express the differences that exist in the
metamorphoses of Insects, and as these terms refer chiefly to the
changes in the outer form, we will here mention them. As already stated,
the locust is, in our own language, said to have an incomplete
metamorphosis, the butterfly a complete one. The term Holometabola has
been proposed for Insects with complete metamorphosis, while the
appellations Ametabola, Hemimetabola, Heterometabola, and
Paurometabola have been invented for the various forms of incomplete,
or rather less complex, metamorphosis. Some writers use the term
Ametabola for Insects that are supposed to exhibit no change of external
form after quitting the egg, the contrasted series of all other Insects being
then called Metabola. Westwood and others use the word Homomorpha
for Insects in which the condition on hatching more or less resembles that
attained at the close of the development, and Heteromorpha for those in
which the form on emergence from the egg differs much from what it
ultimately becomes.
Hypermetamorphosis.
There are certain minute Hymenoptera that deposit their eggs inside the
eggs of other Insects, where the beings hatched from the parasitic eggs
subsequently undergo their development and growth, finding their
sustenance in the yolk or embryo contained in the host-egg. It is evident
that such a life is very anomalous as regards both food and the conditions
for respiration, and we consequently find that these tiny egg-parasites go
through a series of changes of form of a most remarkable character.[84] It
would appear that in these cases the embryonic and post-embryonic
developments are not separated in the same way as they are in other
Insects. We are not aware that any term has yet been proposed for this
very curious kind of Insect development, which, as pointed out by Brauer,
[85] is doubtless of a different nature from the hypermetamorphosis of
Sitaris.
The internal organs for the continuance of the species are known to be
present in a rudimentary stage in the embryo, and it is a rule that they do
not attain their full development until growth has been completed; to this
rule there may possibly be an exception in the case of the Aptera. But
little information of a comparative character exists as to the dorsal vessel
and the changes it undergoes during metamorphosis. There is
considerable difficulty in connexion with the examination of this structure,
but it appears probable that it is one of the organs that changes the least
during the process of metamorphosis.
Lowne informs us that in the imago of the blowfly the great majority of the
hypodermic cells themselves enter into the composition of the chitinous
integument; and it is perhaps not a matter for surprise that the cells
should die on the completion of their functional activity, and should form a
part of the chitinous investment. Some writers say that the chitinous layer
may be shown to be covered by a delicate extima or outer coat.
The number of ecdyses varies greatly in Insects, but has been definitely
ascertained in only a few forms outside the Order Lepidoptera. In
Campodea Grassi says there is a single fragmentary moult, and in many
Hymenoptera the skin that is cast is extremely delicate, and the process
perhaps only occurs twice or three times previous to the pupal stage. In
most Insects, however, ecdysis is a much more important affair, and the
whole of the chitinous integument is cast off entire, even the linings of the
tracheae, and of the alimentary canal and its adjuncts being parted with.
Sir John Lubbock observed twenty-three moults in a May-fly of the genus
Cloëon,[93] this being the maximum yet recorded, though Sommer
states[94] that in Macrotoma plumbea moulting goes on as long as life
lasts, even after the Insect has attained its full size.
Metamorphosis of Blowfly.
We give some figures, taken from Weismann and Graber, of the imaginal
rudiments existing in the larvae of Muscidae. Although by no means
good, they are the best for our purpose we can offer to the reader. Other
figures will be found in Lowne's work on the blowfly now in course of
publication. Weismann's paper[104] is now thirty years old, and, when it
was written, he was not aware of the intimate connexion the rudiments
have with the integument; this has, however, now been demonstrated by
several observers. Pratt states[105] that the formation of the imaginal
discs in Melophagus ovinus takes place in the later stages of the
embryonic development, and after the manner formerly suggested by
Balfour, viz. invagination of the ectoderm.
Fig. 88.—Median longitudinal section through larva of blowfly during the
process of histolysis. (After Graber.) Explanation in text.
Both the regenerative buds and the rudimentary sexual glands are known
to be derived directly from the embryo; neither of them undergoes any
histolysis, so that we have in them embryonic structures which exist in a
quiescent condition during the period in which the larva is growing with
great rapidity, and which when the larva has attained its full growth and is
disintegrating, then appropriate the products of the disintegration so as to
produce the perfect fly.
Our Fig. 88, taken from Graber, represents a longitudinal median section
of a full-grown larva of Musca, in which the processes of metamorphosis
are taking place. The position of some of the more important imaginal
rudiments is shown by it: b1, b2, b3, rudiments of the three pairs of legs of
the imago; an, of antennae; between an and w, rudiment of eye; w, of
wings; h, of halteres; f, fat-body; d, middle of alimentary canal; n, ventral
chain; st, stigma; 6, 7, sixth and seventh body segments.
Physiology of Metamorphosis.
Although the existence of a pupa is to the eye the most striking of the
differences between Insects with perfect and those with imperfect
metamorphosis, yet there is reason for supposing that the pupa and the
pupal period are really of less importance than they at first sight appear to
be. In Fig. 85 we showed how great is the difference in appearance
between the pupa and the imago. The condition that precedes the
appearance of the pupa is, however, really the period of the most
important change. In Fig. 89 we represent the larva and pupa of a bee; it
will be seen that the difference between the two forms is very great, while
the further change that will be required to complete the perfect Insect is
but slight. When the last skin of the larva of a bee or of a beetle is thrown
off, it is, in fact, the imago that is revealed; the form thus displayed,
though colourless and soft, is that of the perfect Insect; what remains to
be done is a little shrinking of some parts and expansion of others, the
development of the colour, the hardening of certain parts. The colour
appears quite gradually and in a regular course, the eyes being usually
the first parts to darken. After the coloration is more or less perfected—
according to the species—a delicate pellicle is shed or rubbed off, and the
bee or beetle assumes its final form, though usually it does not become
active till after a farther period of repose.
Fig. 89.—Larva and pupa of a bee, Xylocopa violacea: A, larva; B, pupa,
ventral aspect; C, pupa, dorsal aspect. (After Lucas.)
CHAPTER VI
Classification.
We have already alluded to the fact that Insects are the most numerous in
species and individuals of all land animals: it is estimated that about
250,000 species have been already described and have had scientific
names given to them, and it is considered that this is probably only about
one-tenth of those that really exist. The classification in a comprehensible
manner of such an enormous number of forms is, it will be readily
understood, a matter of great difficulty. Several methods or schemes have
since the time of Linnaeus been devised for the purpose, but we shall not
trouble the reader to consider them, because most of them have fallen
into disuse and have only a historical interest. Even at present there
exists, however, considerable diversity of opinion on the question of
classification, due in part to the fact that some naturalists take the
structure of the perfect or adult Insect as the basis of their arrangement,
while others prefer to treat the steps or processes by which the structure
is attained, as being of primary importance. To consider the relative
values of these two methods would be beyond our scope, but as in
practice a knowledge of the structures themselves must precede an
inquiry as to the phases of development by which the structures are
reached; and as this latter kind of knowledge has been obtained in the
case of a comparatively small portion of the known forms,—the
embryology and metamorphosis having been investigated in but few
Insects,—it is clear that a classification on the basis of structure is the
only one that can be at present of practical value. We shall therefore for
the purposes of this work make use of an old and simple system, taking
as of primary importance the nature of the organs of flight, and of the
appendages for the introduction of food to the body by the perfect Insect.
We do not attempt to disguise the fact that this method is open to most
serious objections, but we believe that it is nevertheless at present the
most simple and useful one, and is likely to remain such, at any rate as
long as knowledge of development is in process of attainment.
Orders.
The great groups of Insects are called Orders, and of these we recognise
nine, viz. (1) Aptera, (2) Orthoptera, (3) Neuroptera, (4) Hymenoptera, (5)
Coleoptera, (6) Lepidoptera, (7) Diptera, (8) Thysanoptera, (9) Hemiptera.
These names are framed to represent the nature of the wings; and there
is some advantage in having the Orders named in a uniform and
descriptive manner. The system we adopt differs but little from that
proposed by Linnaeus.[107] The great Swedish naturalist did not,
however, recognise the Orders Orthoptera and Thysanoptera; and his
order Aptera was very different from ours.
2. Orthoptera (ὀρθός straight, πτερόν a wing). Four wings are present, the front pair
being coriaceous (leather-like), usually smaller than the other pair, which are of more
delicate texture, and contract in repose after the manner of a fan. Mouth mandibulate.
Metamorphosis slight.
3. Neuroptera (νεῦρον nerve, πτερόν a wing). Four wings of membranous consistency,
frequently with much network; the front pair not much, if at all, harder than the other pair,
the latter with but little or no fanlike action in closing. Mouth mandibulate. Metamorphosis
variable, but rarely slight.
5. Coleoptera (κολεός sheath, πτερόν a wing). Four wings; the upper pair shell-like in
consistency, and forming cases which meet together over the back in an accurate line of
union, so as to entirely lose a winglike appearance, and to conceal the delicate
membranous hind pair. Mouth mandibulate. Metamorphosis great.
6. Lepidoptera (λεπίς scale, πτερόν a wing). Four large wings covered with scales.
Mouth suctorial. Metamorphosis great.
7. Diptera (δίς double, πτερόν a wing). Two membranous wings. Mouth suctorial, but
varying greatly. Metamorphosis very great.
8. Thysanoptera (θύσανος fringe, πτερόν a wing). Four very narrow fringed wings. Mouth
imperfectly suctorial. Metamorphosis slight.
9. Hemiptera (ἡμι half, πτερόν a wing). Four wings; the front pair either leather-like with
more membranous apex, or entirely parchment-like or membranous. Mouth perfectly
suctorial. Metamorphosis usually slight.
We must again ask the reader to bear in mind that numerous exceptions
exist to these characters in most of the great Orders; for instance,
wingless forms are not by any means rare in several of the Orders.
Before remarking further on this system we will briefly sketch two other
arrangements of the Orders of Insects, for which we are indebted to
Packard and Brauer.
Packard's Classification.
Packard has devoted much attention to the subject, and has published
two or three successive schemes, of which the following is the most